View source for Birdwing

{{Short description|Butterflies in swallowtail family}}
{{for|the anatomy of birds|Bird's wing}}
{{more footnotes|date=December 2012}}
{{Paraphyletic group
| auto = yes
| name = Birdwings
| image = Birdwing representatives.jpg
| image_caption = Male (left) and female (right) representatives of ''Trogonoptera'' (top), ''Troides'' (middle), and ''Ornithoptera'' (bottom)
| parent = Troidini
| subdivision_ranks = Genera
| subdivision       =
*''[[Trogonoptera]]'' <small>[[Robert H. F. Rippon|Rippon]], 1890</small>
*''[[Troides]]'' <small>[[Jacob Hübner|Hübner]], 1819</small>
*''[[Ornithoptera]]'' <small>[[Jean Baptiste Boisduval|Boisduval]], 1832</small>
}}

'''Birdwings''' are [[butterflies]] in the [[Papilionidae|swallowtail]] family, that belong to the genera ''Trogonoptera'', ''Troides'', and ''Ornithoptera''. Most recent authorities recognise 36 species, however, this is debated, and some authorities include additional genera. Birdwings are named for their exceptional size, angular wings, and birdlike flight. They are  found across tropical Asia, [[Mainland Southeast Asia|mainland]] and [[Maritime Southeast Asia|archipelagic Southeast Asia]], and [[Australasia]].

Included among the birdwings are some of the largest butterflies in the world: the largest, [[Queen Alexandra's birdwing]]; the second largest, the [[Goliath birdwing]]; the largest butterfly endemic to [[Australia]], the [[Cairns birdwing]]; and the largest butterfly in India, the [[southern birdwing]]. Another well-known species is [[Rajah Brooke's birdwing]], a particularly attractive species and the national butterfly of [[Malaysia]], named after Sir [[James Brooke]], the first [[White Rajahs|White Rajah]] of 19th-century [[Sarawak]].

Due to their size and brightly coloured males, they are popular among collectors of butterflies, but all birdwings are now listed by [[CITES]],<ref name=CITES>CITES (2011). ''[http://www.cites.org/eng/app/appendices.php Appendices I, II and III.]'' Version 27 April 2011.</ref> thereby limiting (and in the case of ''[[Queen Alexandra's birdwing|O. alexandrae]]'' completely banning) international trade.

== Taxonomy ==
=== Genera and species ===
{{col-begin|width=75%}}
{{col-2}}
'''genus:''' ''Troides''

: '''subgenus:''' ''Ripponia''
::* ''[[Troides hypolitus]]'' – Rippon's birdwing
: '''subgenus:''' ''Troides''
:: '''species group:''' ''Troides aeacus''
:::* ''[[Troides aeacus]]'' – golden birdwing
:::* ''[[Troides dohertyi]]'' – Talaud black birdwing
:::* ''[[Troides magellanus]]'' – Magellan birdwing
:::* ''[[Troides minos]]'' – southern birdwing
:::* ''[[Troides plateni]]'' – Dr. Platen's birdwing
:::* ''[[Troides prattorum]]'' – Buru opalescent birdwing
:::* ''[[Troides rhadamantus]]'' – golden birdwing
:: '''species group:''' ''Troides amphrysus''
:::* ''[[Troides amphrysus]]'' – Malay birdwing
:::* ''[[Troides andromache]]'' – Borneo birdwing
:::* ''[[Troides cuneifera]]''
:::* ''[[Troides miranda]]'' – Miranda birdwing
:: '''species group:''' ''Troides haliphron''
:::* ''[[Troides criton]]'' – Criton birdwing
:::* ''[[Troides darsius]]'' – Sri Lankan birdwing
:::* ''[[Troides haliphron]]'' – haliphron birdwing
:::* ''[[Troides plato]]'' – silver birdwing
:::* ''[[Troides riedeli]]'' – Riedel's birdwing
:::* ''[[Troides staudingeri]]''
:::* ''[[Troides vandepolli]]'' – van de Poll's birdwing
:: '''species group:''' ''Troides helena''
:::* ''[[Troides helena]]'' – common birdwing
:::* ''[[Troides oblongomaculatus]]'' – oblong-spotted birdwing

{{col-2}}
'''genus:''' ''Trogonoptera''

:* ''[[Trogonoptera brookiana]]'' – Rajah Brooke's birdwing
:* ''[[Trogonoptera trojana]]'' – Palawan birdwing

'''genus:''' ''Ornithoptera''

: '''subgenus:''' ''Aetheoptera''
::* ''[[Ornithoptera victoriae]]'' – Queen Victoria's birdwing
: '''subgenus:''' ''Ornithoptera''
::* ''[[Ornithoptera aesacus]]'' – Obi Island birdwing
::* ''[[Ornithoptera croesus]]'' – Wallace's golden birdwing
::* ''[[Ornithoptera euphorion]]'' – Cairns birdwing
::* ''[[Ornithoptera priamus]]'' – common green birdwing
::* ''[[Ornithoptera richmondia]]'' – Richmond birdwing
: '''subgenus:''' ''Schoenbergia''
::* ''[[Ornithoptera chimaera]]'' – chimaera birdwing
::* ''[[Ornithoptera goliath]]'' – Goliath birdwing
::* ''[[Ornithoptera meridionalis]]'' – southern tailed birdwing
::* ''[[Ornithoptera paradisea]]'' – paradise birdwing
::* ''[[Ornithoptera rothschildi]]'' – Rothschild's birdwing
::* ''[[Ornithoptera tithonus]]'' – Tithonus birdwing
: '''subgenus:''' ''Straatmana''
::* ''[[Queen Alexandra's birdwing|Ornithoptera alexandrae]]'' – Queen Alexandra's birdwing

{{col-end}}

=== Natural hybrids ===
* [[Troides mixtum|''Troides prattorum'' × ''Troides oblongomaculatus bouruensis'']] — ''Troides mixtum''
* [[Ornithoptera akakeae|''Ornithoptera rothschildi'' × ''Ornithoptera priamus poseidon'']] — ''Ornithoptera akakeae''
* [[Ornithoptera allotei|''Ornithoptera victoriae'' × ''Ornithoptera priamus urvillianus'']] — ''Ornithoptera allotei''

== Description ==

=== Ova ===

After [[mating]], females immediately begin to seek appropriate host plants; climbing [[vine]]s of the genera ''[[Aristolochia]]'' and ''[[Pararistolochia]]'' (both in the family [[Aristolochiaceae]]) are sought exclusively. The female lays her spherical [[egg (biology)|eggs]] under the tips of the vine's leaves, one egg per leaf.

=== Larva ===

[[File:Garudasalabhamlarva.jpg|thumb|right|Early instar larva of ''Troides minos'']]
The [[caterpillar]]s are voracious eaters but move very little; a small group will defoliate an entire vine. If starved due to overcrowding, the caterpillars may resort to [[cannibalism]]. Fleshy spine-like tubercles line the caterpillars' backs, and their bodies are dark red to brown and velvety black. Some species have tubercles of contrasting colours, often red, or pale "saddle" markings. Like other members of their family, birdwing caterpillars possess a retractable organ behind their heads called an ''[[osmeterium]]''. Shaped like the forked [[tongue]] of a [[snake]], the osmeterium excretes a fetid [[terpene]]-based compound and is deployed when the caterpillar is provoked. The caterpillars are also unappealing to most [[predator]]s due to their toxicity: the vines which the caterpillars feed upon contain [[aristolochic acid]], a [[poison]]ous compound known to be [[carcinogen]]ic in [[rat]]s. The feeding caterpillars incorporate and concentrate the aristolochic acid into their tissues, where the poison will persist through [[metamorphosis (biology)|metamorphosis]] and into adulthood.

=== Pupa ===

[[File:Troides.victoriae.pupa.jpg|thumb|right|Pupa of ''Ornithoptera victoriae'']]
Birdwing [[Chrysalis|chrysalids]] are [[camouflage]]d to look like a dead leaf or twig. Before [[pupa]]ting, the caterpillars may wander considerable distances from their host plants. In ''O. alexandrae'', it takes about four months to get from egg to adult. Barring predation, this species can also survive up to three months as an adult.

=== Imago ===

Birdwings inhabit [[rainforest]]s and adults are usually glimpsed along the forest periphery. They feed upon&mdash;and are important long-range [[pollinator]]s of&mdash;nectar-bearing [[flower]]s of the forest canopy, as well as terrestrial flowers, such as [[lantana]]. They are strong flyers and seek sunlit spots in which to bask.

Breeding behaviour varies little between species; the female's role is relatively passive, slowly fluttering from perch to perch while the male performs an elaborate, quivering yet stationary dance 20&ndash;50&nbsp;cm above her.

[[File:Ornithoptera euphorion, Melbourne Zoo.JPG|left|thumb|Copulating pair of ''[[Ornithoptera euphorion]]'' (female above, male below). The sexes are quite dissimilar in appearance, as is typical of species in the genus ''Ornithoptera''.]]
Birdwings are typified by large size (up to a maximum body length of 7.6&nbsp;cm or 3&nbsp;inches and a wingspan of 28&nbsp;cm or 11&nbsp;inches in ''O. alexandrae''), showy colouration (in contrasting shades of green, yellow, black, white, and sometimes blue or orange), and slender, lanceolate forewings. With few exceptions (i.e., the [[New Guinea]]n ''[[Ornithoptera meridionalis|O. meridionalis]]'' and ''[[Ornithoptera paradisea|O. paradisea]]''), the hindwings lack tails. [[Sexual dimorphism]] is strong in ''Ornithoptera'' species only, where males are black combined with bright iridescent green, blue, orange, or yellow while the larger and less colourful females are overall black or dark brownish with white, pale brown, or yellow markings.

Males and females of most ''Troides'' birdwings are similar and have jet black to brown dorsal forewings, often with the [[vein]]s bordered in grey to creamy white. At least one of these darkly-coloured species (''[[Troides rhadamantus|T. rhadamantus]]'') possesses [[thermoreceptor]]s on the anal veins (A2 and A3) of the wings and on the [[antenna (biology)|antennal]] clubs. The antennal receptors of the clubs&mdash;which also possess [[hygroreceptor]]s that measure atmospheric [[humidity]]&mdash;are known as ''sensilla basiconica''. The thermoreceptors are sensitive to sudden increases in temperature; they are thought to help the butterfly [[thermoregulation|thermoregulate]] and avoid overheating while basking.

The colours of most species are [[pigment]]ary (via [[papiliochrome]]); but two species, ''[[Troides magellanus]]'' and the much rarer ''[[Troides prattorum|T. prattorum]]'', are noted for their use of limited-view [[iridescence]]: the yellow of the dorsal hindwings is modified by bright blue-green iridescence which is only seen when the butterfly is viewed at a narrow, oblique angle. This "grazing iridescence" is brought about through [[diffraction]] of [[light]] (after back-reflection) by the wings' extremely steeply-set, multilayered rib-like [[scale (zoology)|scale]]s (rather than the ridge-lamellae of most other iridescent butterflies, such as ''[[Morpho (genus)|Morpho]]'' species). Such limited-view iridescence was previously only known from one other species, the [[Riodinidae|riodinid]] ''[[Ancyluris meliboeus]]''. In ''A. meliboeus'', however, the iridescence is produced by ridge-lamellar scales and features a wider range of colours.

The close evolutionary relationship between ''Troides'' and ''Ornithoptera'' butterflies is well demonstrated by the fact that commercial breeders have produced numerous [[Hybrid (biology)|hybrids]] between the two.

The final and smallest genus is ''Trogonoptera'' with just two species. They resemble each other, being overall black with iridescent green markings and a red head. Females are duller than males.

== Distribution ==

Birdwings are generally found from [[Southeast Asia]] to northern [[Australasia]]. ''Trogonoptera brookiana'' inhabits the [[Thai-Malay Peninsula]], [[Borneo]], [[Natuna Islands|Natuna]], [[Sumatra]], and various surrounding islands.<ref>The World of Birdwing Butterflies. [http://www.nagypal.net/ttbrooki.htm Trogonoptera brookiana.] Accessed 20 May 2012</ref><ref name=arkive>ARKive [https://web.archive.org/web/20060205094236/http://www.arkive.org/species/GES/invertebrates_terrestrial_and_freshwater/Trogonoptera_brookiana/ Rajah Brooke’s birdwing.] Accessed 20 May 2012</ref> ''Trogonoptera trojana'' is [[endemism|endemic]] to [[Palawan]] in the [[Philippines]]. ''Troides'' species are distributed widely across the [[Indomalayan realm]],<ref name="Red Book">{{cite book|last1=Collins |first1=N. Mark |last2=Morris |first2=Michael G. |title=Threatened Swallowtail Butterflies of the World: The IUCN Red Data Book |url=https://www.biodiversitylibrary.org/item/98674#page/7/mode/1up |year=1985 |publisher=[[International Union for Conservation of Nature|IUCN]] |location=Gland & Cambridge |isbn=978-2-88032-603-6 |via=Biodiversity Heritage Library}}</ref> but may be found as far east as [[New Guinea]] in the case of ''[[Troides oblongomaculatus]]''. Some species may be found as far west as [[India]], and are the westernmost distributed of all birdwings. All ''Ornithoptera'' species are found in the northern portion of the [[Australasian realm]], east of [[Max Carl Wilhelm Weber|Weber's line]]; the [[Moluccas]], New Guinea, the [[Solomon Islands (archipelago)|Solomon Islands]], and northeastern [[Australia]].<ref name=Ngypalnet>[http://www.nagypal.net/ttpriamu.htm Ngypalnet]</ref> An outlier is ''[[Ornithoptera richmondia]]'', which may be found in far northeastern [[New South Wales]], [[Australia]] in the southernmost area of its range; the southernmost distribution of all birdwings.{{citation needed|date=July 2014}}

== Status and protection ==

[[File:Ornithoptera alexandrae nash.jpg|thumb|right|''[[Ornithoptera alexandrae|O. alexandrae]]'' has a very small distribution and is the only birdwing on Appendix I of [[CITES]]]]
With the exception of [[Queen Alexandra's birdwing]] (''O. alexandrae''), all birdwings are listed in [[CITES#Appendix II|Appendix II]] of [[CITES]],<ref name=CITES/> and accordingly their trade is restricted in countries that have signed the CITES convention. Exceptions are made for captive-reared specimens, which mainly originate from ranches in [[Papua New Guinea]] and [[Indonesia]].<ref name=CaptiveRearing>United Nations Environment Programme World Conservation Monitoring Centre (2007). [http://ec.europa.eu/environment/cites/pdf/review_butterflies.pdf Review of trade in ranched birdwing butterflies.] European Commission, 2008.</ref> Most species of all three genera have now been reared in captivity, though with significant differences in the quantities reared of each species.<ref name=CaptiveRearing/> ''O. alexandrae'' is listed on [[CITES#Appendix I|Appendix I]]<ref name=CITES/> and therefore cannot legally be traded internationally. At the 2006 meeting of the CITES Animals Committee some suggested ''O. alexandrae'' should be moved to Appendix II, as the conservation benefits of sustainable management perhaps are higher than those of the trade ban.<ref name=CITES2006>CITES (2006). [http://www.iisd.ca/vol21/enb2149e.html Earth Negotiations Bulletin.] {{Webarchive|url=https://web.archive.org/web/20160303182830/http://www.iisd.ca/vol21/enb2149e.html |date=2016-03-03 }} Summary of the 22nd Meeting of the CITES Animals Committee.</ref>

Three ''Troides'' and eight ''Ornithoptera'' species have been given assessments by the [[IUCN Red List]], with classifications ranging from "least concern" to "endangered".<ref name="IUCN">[http://www.iucnredlist.org IUCN Red List] {{webarchive|url=https://web.archive.org/web/20140627094911/http://www.iucnredlist.org/ |date=June 27, 2014}}, entries for ''Troides'' and ''Ornithoptera''.</ref>

[[Richmond birdwing]]s (''O. richmondia'') depend on the plant ''[[Aristolochia praevenosa]]'' which they need for their caterpillars. However, the very similar ''[[Aristolochia elegans]]'' (Dutchman's pipe) which can be found in many Australian backyards, kills the caterpillars.

==Reproduction==

''Ornithoptera'', or the genus of birdwing butterflies, usually reproduce sexually and are [[oviparous]].<ref name="ReferenceA">Straatman, R. Hybridisation of Birdwing Butterflies (Lepidoptera: Papilionidae) In Papua New Guinea. The Lepidopterological Society of Japan. ''Tyô to Ga'' (''Transactions of the Lepidopterological Society of Japan'') Vol. 27, No, 4 1976 156-162.</ref> In butterflies sex is determined by a [[ZW sex-determination system|WW/WZ system]], with a [[heterogametic]] female, reverse of that found in mammals and many other insects, which have a heterogametic male.<ref>Nielsen, J. (2010). A Review of Synandromorphism in the Genus Ornithoptera Boisduval, (Lepidoptera: Papilionidae). Australian Entomologist, 37(3): 105-112.</ref> During copulation males will transfer an ejaculate containing both sperm and accessory substances that can make up to fifteen percent of a males body mass.

==Mating systems==

[[Mating systems]], first explored in evolutionary terms by Darwin, includes all behaviours associated with [[sexual reproduction]].<ref>Emlen ST, Oring LW. (1977). Ecology, sexual selection, and the evolution of mating systems. Science 197:215-223.</ref> Mating systems include all costs and benefits, pre- and postcopulatory competitions, displays and [[mate choice]]. Butterfly mating systems have great variation, including strict monandry, one male and one female, to [[Polyandry in animals|polyandry]], having many mates of the opposite sex. Typically ''Ornithoptera'' tend to be [[Animal sexual behaviour#Polygamy|polygamous]], mating with more than one individual.

===Female choice===

[[Female choice]] can have a serious impact on mate selection and successful reproduction. Several species of ''Ornithoptera'' have been known to create hybrids if they have no access to their own species.<ref name="ReferenceA"/> ''Troides oblongamaculatus'' females have been known to choose to mate with other species such as ''Ornithoptera priamus poseidon'', which will attempt mating if their own species is not to be found near by. The females will typically resist mating attempts by covering their abdomen with their forewings or dropping to the ground, making mating near impossible. Although the females usually resist these mating attempts, they have been noted to be more susceptible if they have not had previous encounters with males of their own species.

===Male courtship===

Some male ''Ornithoptera'' species demonstrate [[courtship display|courtship]] behaviour. ''Ornithoptera priamus posedion'' males will approach a female carefully, and examine the female for several minutes. After consideration, the male may choose to hover twenty to thirty centimeters above the female, displaying the bright yellow marking on its hindwings. Meanwhile, the forewings will move forward, exposing the abdomen and androconial hair tufts. Mating is only attempted when the female has ceased to flap her wings.<ref>Orr, A. (1999). Possible Postcopulatory Mate Guarding in Ornithopter Euphorion (Gray) (Lepidoptera: Papilionidae). Australian Entomologist, 26 (3): 71-76.</ref> After about thirty seconds of the display, the male will attempt copulation.

===Cryptic choice: sperm competition and postcopulatory guarding===

In many animals, females often mate with more than one male.<ref>Satoh, Y. Yano, S. Takafuji, A. Mating strategy of spider mite, Tetranychus urticae (Acari: Tetranychidae) males: postcopulatory guarding to assure paternity. (2001). Applied Entomology, 36 (1): 41-45.</ref> Males who are able will adapt strategies such as postcopulatory guarding to ensure the paternity of the offspring. Following insemination, it is common for the male ''Ornithoptera'' to produce a [[mating plug]], which will seal the ostium bursae and prevent remating by the female, as new sperm is unable to enter the opening. The plug does not impede oviposition and may stay in place for the duration of the female's life.

===Sexual dimorphism===

[[Sexual dimorphism]] is very prominent in ''Ornithoptera'' species, the males being black with brightly colored markings of blue, green, orange or yellow and the females are overall black or dark brown.<ref>Wilts, B. D., Matsushita, A., Arikawa, K., & Stavenga, D. G. (2015). Spectrally tuned structural and pigmentary coloration of birdwing butterfly wing scales. Journal of the Royal Society Interface, 12(111), 20150717. http://doi.org/10.1098/rsif.2015.0717</ref> The sexual dichromatism functions in mate recognition by the use of photoreceptors. Due to the protected nature of ''Ornithoptera'' it has been difficult to study the spectral sensitivities of the sexes although this difference in coloration alludes to the idea of sensory exploitation of the female's photoreceptors. The sensory bias of females to select for males with brighter wings has yet to be studied in ''Ornithoptera''.

Gyanandromorphism is a very rare condition in which an organism simultaneously expresses both male and female phenotypes. It is only observed in species that express strong sexual dimorphism. Gynandromorphs are suspected to be due to genetic errors associated with cell division such as nondisjunction, as well as fertilization of binucleate ova and fertilisation of multiple sperm that may fuse and act as a second nucleus. ''Ornithoptera'' is known to commonly exhibit this phenomenon, but little to no research has been successful in determining why. Those who experience this phenomenon, usually females, show male-pigmented tissues on their wings.

== See also ==

* [[List of largest insects]]

== Cited references ==

<references/>

== Other references ==

{{colbegin}}
* d'Abrera, Bernard. (1975). ''Birdwing Butterflies of the World'' Hill House Publishers {{ISBN|0947352422}}
* American Museum of Natural History. [https://web.archive.org/web/20050817204117/http://sciencebulletins.amnh.org/biobulletin/biobulletin/story845.html BioBulletin: Birdwing butterflies] Retrieved June 28, 2005
* Campbell, A.L., Naik, R.R., Sowards, L., and Stone, M.O. (2002). [https://web.archive.org/web/20030615223813/http://web.neurobio.arizona.edu/gronenberg/nrsc581/thermo/biologicalinfraredsenses.pdf Biological infrared imaging and sensing]. ''Micron 33'', 211&ndash;225.
* Igarashi, S. (1979). Papilionidae and their early stages. Volume I Text (in Japanese), Volume 2 Plates. Kodansha, Tokyo.
* Parsons, M.J. (1996). A phylogenetic reappraisal of the birdwing genus ''Ornithoptera'' (Lepidoptera: Papilionidae: Troidini) and a new theory of its evolution in relation to Gondwanan vicariance biogeography ''[[Journal of Natural History]]'' Volume 30, Issue 11:1707-1736.
* Parsons, M.J. (1996). Gondwanan evolution of the troidine swallowtails (Lepidoptera: Papilionidae): Cladistic reappraisals using mainly immature stage characters, with focus on the birdwings Ornithoptera Boisduval ''Bulletin of the Kitakyushu Museum of Natural History'' 15: 43-118, 34 figures, 2 tables [https://web.archive.org/web/20141011112132/http://www.kmnh.jp/publication/ronbun_pdf/15-43-E-Parsons.pdf pdf]
* Parsons, M.J. (1992). The butterfly farming and trading industry in the Indo-Australian region and its role in tropical forest conservation. ''Tropical Lepidoptera'' 3 (Suppl. 1): 1-31.[http://www.troplep.org/TLR/3supplement1/pdf001.pdf pdf Full text]
* Reed, R.D., and Sperling, F.A.H. (2001). [http://tolweb.org/tree?group=Papilionidae&contgroup=Papilionoidea Tree of Life: Papilionidae] Retrieved June 28, 2005
* {{cite web |last=Savela |first=Markku |date=March 20, 2019 |url=https://www.nic.funet.fi/pub/sci/bio/life/insecta/lepidoptera/ditrysia/papilionoidea/papilionidae/papilioninae/troides/ |title=''Troides'' Hübner, [1819] |website=Lepidoptera and Some Other Life Forms |accessdate=February 4, 2020}}
* Vukusic, P., Sambles, J. R., and Ghiradella, H. (2000). [https://web.archive.org/web/20050829194840/http://newton.ex.ac.uk/research/emag/butterflies/classification_page.htm Optical classification of microstructure in butterfly wing-scales]. ''Photonics Science News'', 6, 66&ndash;66.
* Nagypal, Tony. [http://www.nagypal.net The World of Birdwing Butterflies] {{Webarchive|url=https://web.archive.org/web/20090207013456/http://www.nagypal.net/ |date=2009-02-07 }}.
* Haugum, Jan. (1981). Notes on the ''Aristolochia'' of the Papuan Region, with particular reference to the larval foodplants of the Ornithoptera. ''Lep. Group Newsl.'' 2(10), pp.&nbsp;171–178
* Haugum, Jan; & Low, A. M. (1978). ''A Monograph of the Birdwing Butterflies''. Volume 1, Part 1. Introduction, ''Ornithoptera'' (''Aetheoptera'')., Klampenborg, Denmark, Scandinavian Science Press 1(1)
* Haugum, Jan; & Low, A. M. (1979). ''A Monograph of the Birdwing Butterflies''. Volume 1, Part 2. ''Ornithoptera'' (''Ornithoptera'')., Klampenborg, Denmark, Scandinavian Science Press 1(2)
* Haugum, Jan; & Low, A. M. (1980). ''A Monograph of the Birdwing Butterflies''. Volume 1, Part 3. ''Ornithoptera'' (''Schoenbergia'')., Klampenborg, Denmark, Scandinavian Science Press 1(3)
* Haugum, Jan; & Low, A. M. (1981). ''A Monograph of the Birdwing Butterflies''. Volume 2, Part 1. ''Trogonoptera'' & ''Ripponia''., Klampenborg, Denmark, Scandinavian Science Press 2(1)
* Haugum, Jan; & Low, A. M. (1982). ''A Monograph of the Birdwing Butterflies''. Volume 2, Part 2. ''Troides''; ''amphrysus'' & ''haliphron groups''., Klampenborg, Denmark, Scandinavian Science Press 2(2)
* Haugum, Jan; & Low, A. M. (1983). ''A Monograph of the Birdwing Butterflies''. Volume 2, Part 3. ''Troides''; ''helena'' and ''aeacus'' groups., Klampenborg, Denmark, Scandinavian Science Press 2(3)
* Kiyotaro Kondo, Tsutomu Shinkawa & Hirotaka Matsuka. (2003). Molecular systematics of birdwing butterflies (Papilionidae) inferred from mitochondrial ND5 gene ''Journal of the Lepidopterists' Society'' 57:17-24 [http://peabody.research.yale.edu/jls/pdfs/2000s/2003/2003-57(1)17-Kondo.pdf pdf]{{dead link|date=November 2016 |bot=InternetArchiveBot |fix-attempted=yes}}
* [[Robert Henry Fernando Rippon]] (1898 to 1906) ''Icones Ornithopterorum'' [London] Published by the author at Upper Norwood, London, S.E.
* Schäffler, Oliver . (2001). ''Schmetterlinge der Erde'', ''Butterflies of the World'' Part XII (12), Papilionidae VI: ''Ornithoptera'' Edited by Erich Bauer and Thomas Frankenbach Keltern: Goecke & Evers; Canterbury: Hillside Books. {{ISBN|9783931374839}} Supplement to von Knötgen, 1997
* von Knötgen, Béla. (1997). Ornithoptera: Ornithoptera Schönbergia, Aetheoptera Wangen (Allemagne): MGG Verlag, 1997. Parallel text in German, English and French.
* Darby, A.W. (1982). "The female genitalia of the Birdwing Butterflies, part 1 ''Lepidoptera Group 68''. Vejle. 1982. <small>Showing female genitalia of ''T. helena cerberus, O. priamus richmondia, O. priamus arruana, T. brookiana albescens''</small>.
* Darby, A.W. (1983). "The female genitalia of the Birdwing Butterflies, part 2.  ''Lepidoptera Group'' 68. Vejle. 1983. <small>Showing female genitalia of ''O. goliath procus, T. amphrysus ruficollis, T. a. flavicollis, T. miranda miranda, T. m. neomiranda, T. cuneifera paeninsulae, T. helena cerberus, T. h. hephaestus, T. oblongomaculatus oblongomaculatus, T. o. bouruensis, T. o. papuensis, T. aeacus aeacus, T. a. thomsonii, T. aeacus formosanus, T. rhadamantus rhadamantus, T. r. dohertyi, T. r. plateni, T. vandepolli vandepolli, T. v. honrathiana, T. criton, T. darsius, T. haliphron haliphron, T. h. socrates, T. h. iris, T. h. naias, T. h. pallens, T. prattorum, T. magellanus sonani, T. hypolitus hypolitus''</small>.
{{colend}}

== External links ==
* [https://kagawa-u.repo.nii.ac.jp/records/2000671 Digital Encyclopedia of Birdwing Butterflies – An academic database from Kagawa University featuring high-resolution images and taxonomy of all known species.]
* [http://eol.org/pages/15904/media Images representing ''Ornithoptera''], at EoL
* [https://web.archive.org/web/20080601083415/http://en.butterflycorner.net/Genus-ORNITHOPTERA.937.0.html Butterflycorner.net] (English/German)
* [http://insecta.pro/genus/troides/ "Genus ''Troides''"]. ''Insecta.pro''.
* [http://www.asahi-net.or.jp/~ch2m-nitu/toribaee.htm Birdwings on postage stamps]
* [http://www.pteron-world.com/topics/classfication/papilionidae/troidini.html Pteron] Birdwing Gallery. In Japanese but with [[binomial nomenclature|binomial]] names.

=== Wikimedia sister projects ===
* [[Wikimedia Commons]] has media related to [//commons.wikimedia.org/wiki/Category:Ornithoptera ''Ornithoptera''] and [//commons.wikimedia.org/wiki/Category:Troides ''Troides'']
* [[Wikispecies]] has information related to [//species.wikimedia.org/wiki/Ornithoptera ''Ornithoptera''], [//species.wikimedia.org/wiki/Trogonoptera ''Trogonoptera''], and [//species.wikimedia.org/wiki/Troides ''Troides'']

{{Birdwing |state=expanded}}

{{Taxonbar|from1=Q18086809}}

[[Category:Papilionidae]]
[[Category:Troides|*]]
[[Category:Ornithoptera|*]]
[[Category:Insect common names]]
[[Category:Taxa named by Jean Baptiste Boisduval]]